DEMOGRAPHIC MONITORING OF ARABIS EECUNDA POPULATIONS
IN THE SAPPHIRE AND BEAVERHEAD RANGES, MONTANA
1992 Progress Report
x-icpdrea oy
Peter Leslca s J. Stephen Shelly
Montana Natural Heritage Program
btate Library Building
1515 E. 6th Ave.
Helena, Montana 59620
Prepared for
USDA Forest Service
Beaverhead National Forest
610 N. Montana
Dillon, MT 59725
January 1993
© 1993 Montana Natural Heritage Program
This document should be cited as follows:
Lesica, P. and J. S. Shelly. 1993. Demographic monitoring of
Arabis fecunda populations in the Sapphire and Pioneer Mountains
Ranges, Bearhead National Forest, Montana; 1992 progress report.
20 pp. and appendix.
INTRODUCTION
Passage of the Federal Endangered Species Act of 1973 and
subsequent recognition of the value of conserving biotic
diversity (Wilson 1988) have resulted in many government agencies
becoming active in species conservation. Surveys to determine
the location and size of populations of rare species are being
conducted on public lands throughout the west. These surveys are
necessary in any species conservation program; however, knowing
the location and size of populations at any one point in time is
only the first step in a long-term protection strategy. (Sutter
1986) . Extinction is a process requiring an understanding of
population dynamics (Menges 1986) . Periodic inventories can
detect trends but will do little to determine causality or help
generate predictive hypotheses (Palmer 1987) . Long-term
conservation requires a knowledge of many life history parameters
including fecundity, recruitment, survivorship, age structure,
and population flux. Demographic monitoring techniques can
provide information on factors regulating population density and
persistence (Palmer 1987) . This information, in turn, provides
an essential basis for managemenj: decisions.
Arabis fecunda Rollins is a rosette-forming perennial in the
Mustard Family (Brassicaceae) . This recently described species
(Rollins 1984) is endemic to highly calcareous, azonal soils in
the foothills of the Sapphire Range in Ravalli County and in the
Pioneer and Highland ranges of Beaverhead and Silver Bow
counties, Montana (Lesica 1985, 1992, Schassberger 1988, Lesica
1992) . Arabis fecunda generally occurs on steep, often eroding
slopes with low vascular plant density. In Ravalli County,
populations of A^. fecunda are thought to be threatened by
livestock grazing and encroachment by an aggressive exotic weed,
spotted knapweed ( Centaurea maculosa) (Lesica 1985, Schassberger
1988) . In Silver Bow and Beaverhead counties, populations may be
threatened by mining activity and livestock grazing. Arabis
fecunda is a candidate for listing as a threatened or endangered
species by the U.S. Fish and Wildlife Service (USDI-FWS 1990) and
is considered threatened in Montana (Lesica and Shelly 1991) .
Little is known about the life history and demography of
Arabis fecunda populations. The purpose of this study is to
determine demographic patterns and variability for this rare
species and to use this knowledge to recommend appropriate
management strategies for conservation.
METHODS
Study Sites
We conducted our study at Birch Creek and Charleys Gulch in
Ravalli County and Lime Gulch and Vipond Park in Beaverhead
County, Montana. Both Ravalli County sites are on moderate
1
southwest-facing slopes; the Birch Creek site is 1430 m above sea
level, and the Charleys Gulch site is 1525 m. The two sites are
separated by ca. 10 km. At Hamilton, ca. 8 km southwest and 300
m lower, mean temperatures for July and January are 19.4° and -
3.8° C respectively, and mean annual precipitation is 32 cm.
Zonal vegetation surrounding the sites is foothills Aqropyron-
Festuca grasslands with scattered Pinus ponderosa Dougl. and
Pseudotsuqa menziesii (Mirb.) Franco.
The Lime Gulch site occurs on moderate east- and west-facing
slopes above a small drainage on the east side of the Pioneer
Range at ca 1890 m. The Vipond Park site is on a moderate south¬
facing slope at 2195 m at the north end of the Pioneer Range.
The two sites are separated from each other by ca. 32 km and from
the Ravalli County sites by ca. 130 km. For Divide, at 1675 m
and north and east of the two sites, mean temperatures for July
and January are 17.2° and -7.2° C respectively, and mean annual
precipitation is 31 cm. Zonal vegetation around Lime Gulch is
Juniperus/Cercocarpus woodland, while it is Artemisia-Festuca-
Aqropvron steppe at Vipond Park.
*
Soils at all sites are highly calcareous sandy loams derived
from outcrops of metamorphosed calc-silicates or limestone.
These soils have a tendency to slump on moderate to steep slopes.
Vegetation at these sites is sparse compared to surrounding
grasslands and woodlands. Cryptogamic soil crusts are common at
Birch Creek, Charleys Gulch and Lime Gulch (Lesica and Shelly
1992a) .
Field methods
In 1987 we established two permanent transects, one of 5 and
one of 12 contiguous 1-m^ plots at both Birch Creek and Charleys
Gulch. In 1989 we established two permanent transects of 12
contiguous 1-m^ plots each at both Lime Gulch and Vipond Park.
We censused Arabis fecunda in 1987-91 at Birch Creek, in 1987-
1992 at Charleys Gulch, and 1989-92 at Lime Gulch and’ Vipond
Park. Sampling was conducted in late May at the Ravalli County
sites, mid-June at Lime Gulch and late June or early July at
Vipond Park. We chose these times because ^ fecunda fruits are
mature or nearly so, but dispersal has not yet occurred. Plants
smaller than 0.5 cm in diameter were not recorded because they
could not be reliably distinguished from other species. We did
not distinguish between small- and large-size plants in 1987.
Individual fecunda plants were mapped and recorded using
the following size-life history classification system:
Small (S) = single vegetative rosette < 2 cm in diameter
Juvenile (J) = single vegetative rosette > 2 cm in diameter
Multiple-rosette (M) = multiple vegetative rosettes
Reproductive (R) = plants producing 1-many inflorescences
2
In addition, for each reproductive plant we recorded the number
of inflorescences and the number of fruits matured (see Lesica
1987 for further details) .
A plants 's demographic properties are often more closely
correlated with size and life-history stage rather than age
(Werner and Caswell 1977, Caswell 1989), although both may be
important in predicting an individual's fate (Young 1985). We
chose these classes because they are correlated with age as well
as size and because they also represent a reasonable compromise
between having many categories with too few observations each and
few categories with many observations (Vandermeer 1978) .
In 1989-92 we collected one fruit from the middle of the
inflorescence of each of 25 randomly chosen plants growing near
the transects at each site. We counted the number of mature or
nearly mature seeds in each fruit to obtain an estimate of
seeds/fruit for each site.
We estimated canopy cover of all vascular plants as well as
cover of rock, bare soil and basal vegetation in each plot
(Daubenmire 1959) . We estimated cover to the nearest 5% with two
additional classes, 1-3% and 0-1%.
Data analysis
Population growth for year t is the percent change in the
size of the sample population between year t-l and year t. It is
calculated by PG = Mortality is defined as the
ratio of the number of plants dying between years t-l and t to
the number surviving in the same period. Recruitment rate is
defined as the ratio of new plants observed in year t to the
number of plants surviving from year t-l to year t.
We compared survival rates of uneven-age cohorts present at
the start of the study among the sites using the nonparametric
logrank test (Pyke and Thompson 1986, Hutchings et ai: 1991).
Survivorship curves were constructed following methods outlined
in Hutchings et al. (1991). Probability values were not adjusted
for multiple tests.
For our study, fecundity is defined as the number of fruits
per plant. The effects of site and year on fecundity and number
of seeds/fruit were analyzed using analysis of variance (ANOVA) .
We used ANOVA followed by a contrast test to determine the effect
of site on bare soil in the plots. Plots within transect cannot
be considered independent because they are contiguous. Thus, we
used the eight transects as sampling units for the test.
We used correlation and multiple regression analyses to
explore relationships between weather variables and Arabls
fecunda mortality, recruitment and fecundity. Weather data are
3
from Hamilton and Divide, the two closest recording stations to
the Ravalli County and Beaverhead County sites respectively.
Mean monthly deviation from the 30-year normal for temperature
and precipitation were calculated for the each year up to and
including the spring when demography data were collected (data
from National Oceanic and Atmospheric Administration) . Take 1991
for example, summer included June, July and August, 1990) ; fall
included September, October and November, 1990; winter included
December, 1990 and January and February, 1991; and spring
included March, April and May, 1991. Entries for each site and
each year were used in the analyses. These samples cannot be
considered independent even though responses of A^ fecunda
populations were often very different in the same year (see
Results) . As a result, significance testing of these regressions
is not valid, and reporting probability values is not
appropriate. Nonetheless, these analyses provide useful insight
into environmental factors affecting demography.
RESULTS
Vegetation
Mean canopy cover estimates for dominant vascular plant
species are presented in Table 1. Total basal vegetation cover
was generally similar at the four sites (Table 1) . Graminoids
were common at Lime Gulch, but forbs were more common at the
other three sites. Bare soil was more common at Charleys Gulch
and Lime Gulch, while rock was more abundant at Birch Creek and
Vipond Park. A marginally significant effect on bare soil was
attributable to site (ANOVA; F=4.5, P=0.09), and the amount of
bare soil at Birch Creek and Vipond Park compared together was
greater than at Charleys Gulch and Lime Gulch (P=0.03).
Population Growth
Density of Arabis fecunda at the two Pioneer Range
transects. Lime Gulch and Vipond Park, was 2-3 times higher than
Charleys Gulch or Birch Creek in the Sapphire Range. Over the
course of our study the number of A^ fecunda plants decreased by
ca. 10% at Charleys Gulch and by more than 40% at Birch Creek but
increased by 37% at Lime Gulch and 65% at Vipond Park (Fig. 1) .
Population growth varied among years at all sites (Fig. 2) . The
Charleys Gulch and Lime Gulch populations showed low to moderate
among-year variation and displayed similar patterns in growth
between 1990 and 1992. On the other hand, the Birch Creek and
Vipond Park populations displayed high variability in growth rate
and had similar responses in 1990 and 1991. Responses in growth
rate between Charleys Gulch-Lime Gulch and Birch Creek-Vipond
Park were opposite in sign in three of the five years (Fig. 2) .
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Survivorship
Arabis fecunda mortality rates were more variable and
generally higher at Birch Creek and Vipond Park (Fig. 3) .
Temporal patterns of mortality were more similar between Charleys
Gulch and Lime Gulch and between Birch Creek and Vipond Park.
Survivorship of the uneven-age 1987 cohort declined
significantly more precipitously at Birch Creek than at Charleys
Gulch (log-rank test; LR=19.3, P<0.001), and most of this
difference occurred between 1987 and 1988 (Fig. 4a) . Decline in
survivorship of the uneven-age 1989 cohort was greatest at Vipond
Park and least at Charleys Gulch (Fig. 4b) . Survivorship curves
from Charleys Gulch and Lime Gulch were not significantly
different (LR=1.4, P=0.23), but the Vipond Park survivorship
curve was different from both Lime Gulch (LR=10.2, £=0.001) and
Charleys Gulch (LR=9.5, P=0.001). At Charleys Gulch and Lime
Gulch the probability of death was nearly constant, while at
Birch Creek and Vipond Park younger plants had a higher chance of
dying.
Recruitment
Recruitment of Arabis fecunda varied among sites and years
but was generally higher and more variable at Birch Creek and
Vipond Park (Fig. 5) . The best years for recruitment were not
the same among the four sites. Recruitment was high at Charleys
Gulch and Lime Gulch in 1991 but not at Birch Creek or Vipond
Park. 1988 was a strong recruitment year at Birch Creek but not
Charleys Gulch, and 1992 was the highest recruitment year at
Vipond Park but was lowest for both Charleys Gulch and Lime
Gulch. Recruitment was high at all sites in 1990 (Fig 5.).
There was a strong negative correlation between mortality rate
and recruitment rate among the four sites over the six years of
the study (N=15, r=.63, P=0.01).
Fecundity
The percent of each Arabis fecunda populations that flowered
and produced fruit varied among years (Fig. 6) . On years of high
recruitment a larger percentage of plants are new recruits and
less likely to flower. Thus, the percentage of plants flowering
is expected to be lower in years of high recruitment; however,
this does not appear to be the case (compare Fig. 5 and 6) .
Percent reproductive plants was at least 15% at all sites in all
years except there were few reproductives at Birch Creek and
Charleys Gulch in 1988 and at Lime Gulch in 1989 (Fig. 6) .
Percent reproductives was highest in 1990 at all sites except
Birch Creek.
The number of fruits per reproductive A^ fecunda plant
varied greatly among sites and years (Fig. 7) . Site was a
7
Percent Mortality
Figure 3. Mortality rate of Arabia fecunda populations at four study sites in
1987-1992 (see Methods for definition of mortality rate).
1988 1989 1990 1991 1992
Year
8
Figure 5. Recruitment rate of Arabia fecunda populations at four study sites
in 1987-1992 (see Methods for definition of recruitment rate).
PI
P
1
III
m
1988 1989 1990 1991 1992
^^1
Figure 6. Percent of Arabia fecunda plants that produced flowering
inflorescences at four study sites in 1987-1992.
Charleys
Vipond
1987 1988 1989 1990 1991
significant source of variation in fecundity (ANOVA; N=1461,
F=25.7, P<0.001). None of the sites were significantly different
from any others as determined by contrast test after the ANOVA
(P>0.15), but Birch Creek and Vipond Park together, the two sites
with the greatest overall fecundity, were significantly higher
than Charleys Gulch and Lime Gulch (P<0.001).
The number of seeds per fruit varied from 5 to 56, and means
varied from 26 {SE=2.0) at vipond Park in 1990 to 38 (SE=1.1) at
Birch Creek in 1987 (Table 2). The number of seeds per fruit
differed significantly among years (ANOVA; N=325, F=9 . 3 , P<0.001)
but did not differ significantly among sites (F=1.4, P=0.25) .
Weather and life history
Mortality was negatively associated (N=15, r=0.61) and
recruitment was positively associated (N=15, r=0.48) with winter
precipitation. Higher fecundity (fruits/plant) was associated
with warmer fall and winter temperatures and higher fall
precipitation (multiple regression r=0.78, N=19) .
Overall, there were few consistent relationships between
weather and Arabis f ecunda demographic parameters. The behavior
of populations at different sites was highly idiosyncratic. For
instance, mortality was well below normal at Birch Creek, Lime
Gulch and Vipond Park in 1989-90 but was above average at
Charleys Gulch. Recruitment was higher than average at Charleys
Gulch and Lime Gulch in 1990-91 but was below normal at Birch
Creek and Vipond Park. Fecundity was high at Birch Creek and
Charleys Gulch in 1989 but was below average at Lime Gulch and
Vipond Park.
DISCUSSION
Arabis f ecunda appears to have two distinctive life-history
strategies. Populations at Charleys Gulch and Lime Gulch showed
similar demographic responses and had higher survivorship, lower
mortality, lower recruitment rates and lower fecundity compared
to populations at Birch Creek or Vipond Park. Arabis fecunda
populations at Charleys Gulch and Lime Gulch had similar
demography and were less variable in size and growth rate as well
as demonstrating less variation in recruitment and mortality
rates than those at Birch Creek or Vipond Park. Populations at
Birch Creek and Vipond Park have a high variation strategy, while
those at Charleys Gulch and Lime Gulch have a more stable state
strategy.
Intraspecific differences in demography can be the result of
genetic differentiation among populations and/or phenotypic
plasticity. The existence of intraspecific ecotypes that differ
12
Table 2 . Number
sites in 1989-91
Birch Creek
Charleys Gulch
Iiime Gulch
Vipond Park
of seeds per fruit (+SE) for Arabia fecunda at the four study
1989
38.6 (1.1)
32.3 (0.7)
31.1 (1.1)
35.4 (1.6)
1990
30.4 (1.1)
31.2 (1.0)
34.3 (1.5)
26.0 (2.1)
1991
32.1 (1.1)
31.8 (0.9)
33.8 (1.5)
35.7 (2.0)
13
in demography has long been recognized (Turesson 1922, Clausen et
al. 1940, Stebbins 1950) , and different ecotypes can occur within
short distances of each other (Solbrig and Simpson 1977) . Under
selection pressure from the environment, different populations
can evolve divergent genotypes highly adapted to their
surroundings. On the other hand an "all-purpose genotype" adapts
to different environments by plastic responses in morphology,
phenology and demography (Bradshaw 1965, Schlichting 1986) .
These two means of coping with environmental variability are not
mutually exclusive, and either or both could be the source of
variable demographic strategies in Arabis fecunda.
Regardless of the underlying mechanism causing demographic
differentiation in Arabis fecunda . it is interesting to speculate
on the environmental factors that drive the differentiation. The
Birch Creek and Charleys Gulch populations occur at 1430-1530 m
in elevation and are less than 10 km apart in the Sapphire Range.
They are found in similar soils derived from the same geologic
formation * Lime Gulch and Vipond Park are at 1890 and 2195
respectively and are ca. 30 km apart in the Pioneer Range. The
two sites in the Pioneer Range ai;e 13 0 km south of the two
Sapphire Range sites. If abiotic factors such as weather or
soils were determining life history strategies, we would expect
there to be more similarity between the more geographically
proximal sites, but this is not the case. The fact that the
Birch Creek population is demographically more similar to Vipond
Park than Charleys Gulch and that Charleys Gulch is more similar
to Lime Gulch than Birch Creek indicates that abiotic factors,
meteorological or edaphic, are not responsible for the
demographic differentiation. Arabis fecunda habitats at Birch
Creek and Vipond park have, on average, 15% more bare soil than
those at Charleys Gulch and Lime Gulch, while the total cover of
vascular plants is similar. Thus, ^ fecunda populations that
experience greater interspecific crowding are also the ones with
higher and more variable demographic parameters.
Crowding has been shown to increase mortality of younger or
smaller plants (Harper 1977, Powell 1990), Competition for water
or nutrients could increase variability in mortality rate by
amplifying the effects of adverse weather conditions. For
example, a crowded population would be expected to suffer heavy
mortality in a dry year, while a population that is not competing
for water may not show such a large decrease in survivorship. In
wet years, mortality at the two sites would be more similar.
Recruitment of A^. fecunda is enhanced by reduced interspecific
competition (Lesica and Shelly 1992b) and should be lower in less
crowded habitats. However, if differences between the two
demographic strategies are under genetic control, increased
fecundity and recruitment may have evolved to compensate for
increased mortality in crowded habitats. Experimental work is
needed to determine the basis for the two demographic strategies
found in A^. fecunda populations. Nonetheless, our results suggest
14
that there may be genetic differentiation for demographic
parameters among populations of fecunda.
In many instances populations of Arabis fecunda at Birch
Creek and Vipond Park had a different response than the other two
populations to similar weather. In spite of this, our results
suggest that winter precipitation is an important determinant of
survivorship and recruitment. Low mortality often occurred in
years with wet winters, and low recruitment often occurred
following dry winters. The importance of winter precipitation
could be in providing a protective blanket of snow during the
coldest months or in providing ample early-season moisture. The
importance of winter precipitation is probably at least partially
responsible for the strong negative correlation between mortality
and recruitment. There was no correlation between fecundity and
recruitment or between fecundity and mortality (N=15, r<0.08).
Higher fecundity was associated with above normal fall and winter
temperatures and above normal fall precipitation. Since even in
a poor year a reproductive plant produces an average of at least
120 seeds, it seems unlikely that year-to-year fluctuations in
fecundity would have an important effect on population growth.
Thus, it appears that winter precipitation is the most important
weather variable affecting ^ fecunda population growth.
MANAGEMENT IMPLICATIONS
A recent study of Arabis fecunda using isozyme markers
suggests that there is little genetic differentiation among
populations (Leeper et al. 1993). However, isozyme studies
detect differentiation of neutral genotypes that occurs over
relatively long periods of time. Genetic distance based on
isozyme markers is generally related to length of time since
population separation and degree of gene flow. Significant
selection-driven differentiation can occur without
differentiation among neutral alleles. Our study suggests the
existence of ecotypic differentiation among ^ fecunda
populations. If differentiation does exist, a conservation
strategy for this rare state endemic must strive to protect the
genetic variation present among the many populations.
The Birch Creek and Vipond Park populations of Arabis
fecunda demonstrated relatively large fluctuations in population
size compared to those at Charleys Gulch and Lime Gulch. For
example, population growth decreased in all populations in 1991;
however, the decrease was less than 5% at Charleys Gulch and Lime
Gulch but was 50-90% at Birch Creek and Vipond Park (Fig. 2).
Populations subject to large fluctuations are generally thought
to be more prone to extinction (Diamond 1984, Pimm et al. 1988).
The Birch Creek and Vipond Park populations are large
(Schassberger 19??) , but smaller populations with the same
demographic behavior may be at risk if prolonged stress occurs.
15
There is widespread agreement that increasing levels of
atmospheric carbon dioxide and other "greenhouse gases" will
result in global warming and widespread alteration of rainfall
patterns during the next 50 years (Peters 1988, Henderson-Sellers
1990) . Drier conditions are expected to cause a decrease in
recruitment and an increase in mortality in Arabis fecunda
populations. Warmer winters may result in some increase in
fecundity and may partially offset the effects of drier
conditions on mortality and recruitment. Nonetheless, if winter
precipitation in southwest Montana decreases, populations of A.
fecunda may decline. The greenhouse effect is also predicted to
cause an increase in the variation of weather events (Schneider
et al. 1992) . Increased environmental variation could interact
with the high demographic variation of some A^ fecunda
populations to increase the chances of extinction.
In summary, many populations of Arabis fecunda from a
variety of habitats should be given protection to ensure
maintenance of genetic diversity. Small populations that
demonstrate widely fluctuating demographic behavior are less
appropriate for long-term conservation purposes. Populations in
relatively mesic habitats are expected to have longer persistence
and should be given priority over those in drier sites.
ACKNOWLEDGEMENTS
Lisa Schassberger Roe, Peter Achuff and Bonnie Heidel helped
conduct field work. Portions of this study were funded by
Beaverhead National Forest, the U.S. Fish and Wildlife Service
and The Nature Conservancy.
16
LITERATURE CITED
Bradshaw, A. D. 1965. Evolutionary significance of phenotypic plasticity in
plants. Advances in Genetics IBs 115-155.
Caswell, H. 1989. Matrix population models. Sinauer Associates, Sunderland,
Massachusetts, USA.
Clausen, J., D. D. Keck and W. M. Hiesey. 1940. Experimental studies on the
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17
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18
Appendix A. Performance of individual Arabis fecunda plants in
permanent monitoring transects on Beaverhead National Forest. See
Methods for description of codes. Individuals are identified by
letters within each plot. A "b" indicates plants that
have''bolted, " produced a large, multi-branched inflorescence from
the terminal bud.
Vipond Park West
Plot
1989
1990
1991
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—
—
—
R1
s
—
--
—
S
t
—
—
—
R1
u
—
—
—
R1
V
—
—
—
R1-I1-F2
w
—
—
—
S
X
—
—
— —
R1-I1-F7
9 .
R1-I2-F7
—
—
—
b
S
R1
R1
R1-I3-F11
c
R1-I4-F26
--
—
—
d
R1
R1-I2-F6
R1
R1-I3-F8
e
R1
Rl-Il-F23b
—
—
f
R1-I2-F11
Rl-Il-F30b
—
g
R1
R1
R1
R1-I4-F15
h
R1
—
—
—
i
R1-I1-F3
R1-I1-F2
—
—
j
R1
—
—
—
k
R2
R2
R2
R2-I1-F4
1
R2
—
—
m
R1-I1-F4
R2-Il-F27b
R1
R1-I3-F7
n
R1
R1
— ■
--
o
—
R1-I2-F8
—
—
P
—
R1
—
—
q
—
S
R1
—
r
—
S
R1
—
s
—
R1
R1-I1-F4
R2-I5-F15
t
—
S2
—
—
u
—
R1
R1-I1-F7
Rl-Il-F27b
V
—
R1
—
--
w
—
R1-I5-F16
R1-I1-F7
Rl-Il-F27b
X
—
R1
R1
R1-I2-F7
y
—
R2
R2
R2-I3-F22b
z
—
S
SI
--
aa
—
Rl-Il-F53b
—
—
bb
—
—
R1
R1-I5-F15
cc
--
—
S
S2
dd
—
—
R1
S2-Il-F5b
ee
—
—
S
ff
—
—
S
gg
—
--
—
S
hh
—
—
—
S
ii
—
—
—
S2
jj
—
—
—
R1
kk
—
--
—
R1
11
—
—
—
R1
10.
R1
R1
R1-I2-F6
—
b
R3-I1-F9
—
—
—
c
R3
R2-I3-F5
—
—
d
R3-I1-F1
R2-I5-Fllb
R1-I1-F2
—
e
R2
R2
R2
R3-Il-F26b
f
R1
S
—
—
g
R1
Rl-Il-F27b
—
—
h
R1
R1-I1-F4
R1
R1
i
R3
R2
R1
R1
j
—
S2
—
--
k
—
S
S
S2
1
—
S
S
R1
m
—
S
R1
—
n
—
—
R2-Il-F8b
S2
o
—
—
S
—
P
—
—
—
S
q
—
—
—
Rl-Il-F21b
r
—
—
—
S3
11.
R1
RlSl
Rl-Il-F8b
Rl-Il-F15b
b
R1
R1-I3-F8
R1-I2-F12
Rl-Il-F21b
c
S3
R1S2
R5
R3-I4-F28b
d
R2-I3-F13
—
—
—
e
R1
R1-I1-F3
R1-I2-F11
R1-I2-F7
f
R1
Rl-Il-F15b
—
—
g
S
Rl-Il-FlO
—
—
h
—
Rl-Il-F32b
—
—
i
— -
R1S2
R3
R1S2-I2-F2:
j
—
R1
R1
R1-I2-F4
k
—
Rl-Il-F23b
—
—
1
—
S2-I2-F9b
—
—
m
—
RlSl
R5
R5
n
—
—
Rl-Il-F14b
Rl-Il-F12b
o
—
—
S
R1
P
—
—
—
S2
q
—
—
—
SI
r
—
—
—
R1
12.
s
R1-I1-F3
—
—
b
R3
—
—
—
c
R1
R1-I4-F22
Rl-I2-F21b
--
d
R1-I1-F4
R1-I2-F2
R1-I4-F36
Rl-Il-F23b
e
R1
R1-I1-F2
—
—
f
R1
R1
R1-I1-F6
R1-I1-F3
g
S
R1
S2
S3
h
S
—
—
—
i
S2
—
—
j
S
R1
—
—
k
S
S
R1
R1-I1-F2
1
R1
Rl-Il-F27b
—
—
m
R1
R1
R1
R1
n
R1
R1-I1-F5
R1
—
o
R1
Rl-Il-F52b
—
—
P
R1
Rl-Il-F19b
—
—
q
R1-I3-F11
Rl-Il-F46b
—
—
r
R4-I1-F3
R3-Il-F24b
R2-I1-F7
Rl-Il-F29b
s
R1
Rl-Il-F37b
—
—
t
R1
Rl-Il-F50b
—
—
u
R1
R1-I1-F3
—
—
V
—
R1
R1
R1
w
—
—
R1
R1
X
—
—
S
—
y
—
—
S
S
z
—
—
R1
R1
aa
—
—
S
S2
bb
—
—
S3
—
cc
—
—
S
—
dd
—
—
R1
R1-I3-F13
ee
—
—
S
R1
ff
—
—
S
S
gg
—
—
R1
R2
hh
—
—
—
S
ii
—
—
—
R1-I1-F5
j j
—
—
—
R1
kk
—
—
—
S
11
—
—
—
S
mm
—
—
R1-I1-F4
nn
—
—
—
S
oo
—
—
—
S
pp
—
—
—
S
qq
— —
— —
s
rr
Vipond
Park East
R1
Plot
1989
1990
1991
1992
1.
R1
R1
Rl-Il-F25b
—
b
S
R1-I1-F4
R1-I5-F19
—
c
R1-I2-F5
Rl-Il-F45b
—
—
d
R2
R2-I2-F54b
—
—
e
R3
—
—
—
f
R2
RlSl
R2-I1-F4
R2-I3-F18b
g
R2
—
—
—
h
R1-I2-F8
—
—
—
i
R1
Rl-Il-F50b
—
—
j
S
R1
R1
R1-I3-F10
k
s
R1
RlSl
RlSl-Il-F13b
1
—
S2
—
—
m
—
Rl-Il-F19b
—
—
n
—
S
' S
S2
o
—
R1
R1
R1
P
—
S
R1
R1
q
—
R1
R1
R1-I1-F7
r
--
R1-I2-F15
R1
--
s
—
R1
R1
R1-I1-F24
t
—
R1
R1-I1-F3
R1
u
—
R1
—
—
V
—
Rl-Il-F20b
—
--
w
—
R1
R2
R2-I3-F10
X
—
S2
R1
R1-I5-F13
y
--
R1-I3-F12
—
—
z
—
R1
R1
R1
aa
—
R1
R1
R1
bb
—
—
S
S2
cc
—
—
R1
— .
dd
—
—
R1
R1
ee
—
—
S
—
ff
—
—
S2
R1
gg
—
—
R1
—
hh
—
—
R1
S
ii
—
—
S
S
j j
—
—
R1
R1-I1-F3
kk
—
—
R1
R1
11
—
—
R2
S
mm
—
—
R1
Rl-Il-FlO
nn
— ■
—
—
R1
oo
—
—
—
R1
PP
—
—
—
R1
qq
—
—
—
R1
rr
—
—
—
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ss
—
— -
—
R1-I3-F8
tt
—
■ —
—
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uu
—
—
—
S
w
—
—
—
S2
ww
--
—
—
S
XX
—
—
—
S
yy
— —
—
S
2.
R1
Rl-Il-F21b
—
—
b
R1
R2
R3-I2-F21b
—
c
S
—
—
—
d
R1
R1-I1-F2
Rl-Il-FlO
RlSl-Il-F32b
e
S
R1
R1
—
f
R3-I3-F20
R3-I1-F3
S2
—
g
R1
R1-I1-F2
S2
R2
h
R2-I2-F4
R2-I2-F40b
—
—
i
—
S1-I1-F4
—
—
j
—
R1
R1
—
k
—
R1
RlSl
—
1
—
S
R1
—
m
, —
S
S3
S3
n
—
Rl-Il-F9b
—
—
o
—
S
R4-I1-F5
—
P
—
R1
—
—
q
—
R1
R1
R3
r
—
R1
—
—
s
—
S
—
--
t
—
R1
—
—
u
—
R1
R1
SI
V
—
S
R1
—
w
—
R1
—
—
X
—
R1-I5-F25
—
—
y
—
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S
s
z
—
S
S
S7
aa
—
R1
R1
—
bb
—
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R1
Rl-Il-F16b
cc
--
S
S2
--
dd
—
—
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—
ee
—
—
R1
R1
ff
—
—
R1
R2
gg
—
—
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— .
hh
—
—
S2-I1-F3
RlSl-Il-FOb
ii
— ■ ■
—
S1-I1-F4
—
j j
—
—
S
S2
kk
—
—
S
S
11
—
—
s
RlSl-Il-Fl
mm
—
—
R1
R2
nn
—
—
S
R3-I2-F17b
oo
—
—
R1
Rl-Il-F9b
PP
—
—
S
—
qq
— —
—
S
— —
rr
—
—
R1
R1
ss
—
—
—
RlS2-Il-F12b
tt
—
—
—
S
uu
—
—
—
R1
w
—
—
—
R1-I1-F3
ww
—
—
—
S
XX
- 7
—
—
R1
yy
—
—
—
S
zz
—
—
—
R1
aaa
—
—
—
R1
bbb
—
—
—
S
ccc
—
—
—
S2
ddd
—
—
—
S
eee
— —
— —
—
R1
3 .
R1-I1-F3
—
—
—
b
R1
—
—
—
c
R1-I1-F3
Rl-Il-F32b
--
—
d
R1-I3-F13
Rl-Il-F14b
—
—
e
R1
—
—
—
f
S
Rl-Il-F38b
—
—
g
R1-I1-F5
Rl-Il-F29b
—
—
h
R1-I4-F16
—
—
—
i
R1-I1-F3
S
S
—
j
R1-I2-F9
S
Rl-Il-F51b
—
k
S2
R2-I4-F29b ,
—
1
R1
Rl-Il-F9b
—
—
m
R1
R1-I6-F33
—
—
n
R1
Rl-Il-FlOb
—
—
o
R1-I2-F4
Rl-Il-F24b
— ‘
—
P
—
S
—
—
q
—
Rl-Il-F19b
—
—
r
—
Rl-Il-F27b
—
—
s
—
S
R1
R1
t
—
—
S
R1
u
--
—
S
S
V
—
—
R1
RlSl
w
—
—
R2-I3-F14
—
X
—
—
S
S
y
—
—
S
s
z
—
—
R1
R1
aa
—
—
S
S4
bb
—
—
—
Rl-Il
cc
—
—
—
S
dd
—
—
—
S2
ee
—
—
■ —
S
ff
■ —
—
—
S
gg
—
—
—
S
hh
—
—
—
S
ii
—
—
—
S
j j
—
—
—
S
kk
—
—
—
s
11
—
—
--
s
mm
—
—
—
s
nn
—
—
—
s
oo
—
—
—
s
PP
—
—
—
R1
qq
— —
— —
— —
S
rr
—
—
—
s
1 ss
S
tt
—
—
—
s
uu
--
—
—
s
5 W
—
—
—
s
J ww
—
—
—
s
XX
— —
—
—
s
"1 4 .
R1
R1
—
—
1 b
R4
R3-I3-F18b
—
—
c
R3-I1-F1
R4-I4-F21b
—
—
1 d
R1
R1
R1-I1-F9
Rl-Il-F28b
1 ' e
R1
—
—
—
f
R1-I1-F2
—
—
—
g
—
R3
R3
R3
i' h
—
S
—
—
i i
—
S
—
—
j
—
S2
S
—
1 k
—
S
—
—
1 ^
—
R1-I2-F2
S4
S2
m
—
S
—
—
i n
—
R1
R3-I2-F11
R3
1 °
—
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S
S
i p
—
—
S
S3
q
—
—
R1
—
I ^
—
—
S
S
i s
—
—
s
—
t
—
—
R1
R1-I1-F2
—
—
—
S
1
—
—
—
R2
w
—
—
—
S
X
—
—
—
R1
1 y
—
—
—
S
1 z
—
—
—
S
aa
—
—
—
S
1 bb
—
—
—
S
1 cc
—
--
—
S
dd
—
—
—
S
ee
—
—
—
R1
i ■ ff
—
—
—
S
^ gg
—
—
—
S
hh
—
—
—
S
; ii
—
—
—
S
! j j
—
—
—
S
kk
—
—
—
s
1 11
—
—
—
S6-I5-F13b
1 mm
--
—
—
—
nn
—
—
—
S
oo
—
—
—
s
I PP
—
—
—
s
qq
—
—
— —
s
rr
—
— —
—
s
ss
--
—
—
s
tt
—
—
—
s
uu
—
—
—
s
w
—
—
—
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ww
—
—
—
S
XX
—
—
—
S
yy
—
—
—
S
zz
*—
——
— —
S
5.
R1
R1
R2-I2-F14
—
b
R1
R1-I4-F18
R1-I1-F7
Rl-Il-F32b
c
R1
—
—
d
R1
R1
R1
R1
e
S
R1
RlSl
—
f
S
—
—
—
g
S
R1
R1
R1-I1-F6
h
R2
R2-I1-F5
R2-I1-F3
R2-I2-F16b
i
S
S
S
R1
j
S
—
—
—
k
S
Q
R1
R1
—
X
m
O
s
Rl-Il-F8b
—
—
n
R1-I1-F25
—
—
—
o
R1
R1
' R1
Rl-Il-F7b
P
R1-I4-F9
Rl-Il-F21b
—
—
q
—
S
R3
—
r
--
R1
—
—
s
—
S3
RlSl
—
t
—
S
S
—
u
—
R1-I2-F11
—
—
V
R1
R1-I2-F8
R1-I2-F5
w
—
S
— .
—
X
—
—
R1
--
y
—
—
S2
—
z
—
—
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S2
aa
—
—
—
S
bb
—
—
—
R1
cc
—
—
—
S
dd
—
—
—
S
ee
‘ —
—
—
R1-I2-F4
6.
R2
R1-I1-F6
R1
R1S2
b
S
S
—
—
c
R2
R2
—
—
d
R1
Rl-Il-F8b
—
—
e
R2
R2-I5-F31b
—
—
f
R1
R2
R3
Rl-I3-F23b
g
R1
—
—
—
h
R4-I1-F1
S
R1
Rl-Il-Fl
i
R1-I4-F19
Rl-Il-F26b
—
—
j
R2
R4-I4-F74b
R1
R1
k
R1
R1
R2
1
R1
—
—
— ■
m
R1
R1
Rl-Il-F41b
—
n
R2-I3-F15
R2-I6-F21b
- —
—
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R1-I1-F13
R1S3
R4
—
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— .
R1
R1
R1
q
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S
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1 r
—
R1
R1-I4-F21
R1
s
--
R1-I7-F43
R1-I2-F12
—
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—
—
R1
—
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—
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R1
—
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—
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S3
S3
w
—
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—
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— —
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— —
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R1
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—
—
—
Rl-Il-F20b
7.
R1-I1-F4
R2-I2-F17b
- —
—
1 ^
R1-I1-F9
Rl-Il-F42b
—
—
1 c
R1
R1-I2-F2
Rl-Il-F74b
—
d
R1-I3-F14
R3-I4-F28b
—
—
1 e
R1
—
—
—
I f
S4
R5
—
—
g
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—
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R1
R1-I1-F4
—
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R3
R4-I2-F4
R3-I1-F11
R3-I8-F56b
j
S2
R2
R2
R2-I4-F14
k
R1-I2-F8
Rl-Il-F19b
—
—
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R1
—
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R1
R1-I1-F3
—
—
n
R1
R1-I1-F3
—
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—
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1987 1988 1989 1990 1991 1992
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